Dhole

The etymology of "dhole" is unclear. The possible earliest written use of the word in English occurred in 1808 by soldier Thomas Williamson, who encountered the animal in Ramghur district, India. He stated that dhole was a common local name for the species. In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'. Two years later, Smith connected this word with 'mad, crazy', and erroneously compared the Turkish word with and (cfr. also ; ), which are in fact from the Proto-Germanic *dwalaz 'foolish, stupid'. Richard Lydekker wrote nearly 80 years later that the word was not used by the natives living within the species' range. The Merriam-Webster Dictionary theorises that it may have come from the . Other English names for the species include Asian wild dog, Asiatic wild dog, Indian wild dog, and red wolf. ==Taxonomy and evolution==

dating back to the upper Würm period from Cova Negra de Xàtiva, Valencia, Spain , one of the first accurate depictions of the species, based on a single skin purchased in the village of Dshare on the Amur is not relevant for taxonomy--> Canis alpinus was the binomial name proposed by Peter Simon Pallas in 1811, who described its range as encompassing the upper levels of Udskoi Ostrog in Amurland, towards the eastern side and in the region of the upper Lena River, around the Yenisei River and occasionally crossing into China. This northern Russian range reported by Pallas during the 18th and 19th centuries is "considerably north" of where this species occurs today. Hodgson later took note of the dhole's physical distinctiveness from the genus Canis and proposed the genus Cuon. The first study on the origins of the species was conducted by paleontologist Erich Thenius, who concluded in 1955 that the dhole was a post-Pleistocene descendant of a golden jackal-like ancestor. The paleontologist Bjorn Kurten wrote in his 1968 book Pleistocene Mammals of Europe that the primitive dhole Canis majori Del Campana 1913 —the remains of which have been found in Villafranchian era Valdarno, Italy and in China—was almost indistinguishable from the genus Canis. In comparison, the modern species has greatly reduced molars and the cusps have developed into sharply trenchant points. During the Early Middle Pleistocene there arose both Canis majori stehlini that was the size of a large wolf, and the early dhole Canis alpinus Pallas 1811 which first appeared at Hundsheim and Mosbach in Germany. In the Late Pleistocene era the European dhole (C. a. europaeus) was modern-looking and the transformation of the lower molar into a single cusped, slicing tooth had been completed; however, its size was comparable with that of a wolf. This subspecies became extinct in Europe at the end of the late Würm period, but the species as a whole still inhabits a large area of Asia. and what is believed to be dhole remains have been found at Riparo Fredian in northern Italy dated 10,800 years old. The vast Pleistocene range of this species also included numerous islands in Asia that this species no longer inhabits, such as Sri Lanka, Borneo and possibly Palawan in the Philippines. Middle Pleistocene dhole fossils have also been found in the Matsukae Cave in northern Kyushu Island in western Japan and in the Lower Kuzuu fauna in Tochigi Prefecture in Honshu Island, east Japan. Dhole fossils from the Late Pleistocene dated to about 10,700 years before present are known from the Luobi Cave or Luobi-Dong cave in Hainan Island in south China where they no longer exist. Additionally, fossils of canidae possibly belonging to dhole have been excavated from Dajia River in Taichung County, Taiwan. A single record of the dhole is known from North America. This consists of a jaw fragment and teeth of Late Pleistocene age found in San Josecito Cave in northeast Mexico, dating to around 27,000–11,000 years ago. Other researchers have either considered this record as "insufficient" or suggested that further corroboration is required for the definitive taxonomic attribution of these specimens. }} Dholes are also known from the Middle and Late Pleistocene fossil record of Europe. In 2021, the analyses of the mitochondrial genomes extracted from the fossil remains of two extinct European dhole specimens from the Jáchymka cave, Czech Republic dated 35,000–45,000 years old indicate that these were genetically basal to modern dholes and possessed much greater genetic diversity. Subsequent authors, including Juliet Clutton-Brock, noted greater morphological similarities to canids of the genera Canis, Dusicyon and Alopex than to either Speothos or Lycaon, with any resemblance to the latter two being due to convergent evolution. Some authors consider the extinct Canis subgenus Xenocyon as ancestral to both the genus Lycaon and the genus Cuon. This closeness to Canis may have been confirmed in a menagerie in Madras, where according to zoologist Reginald Innes Pocock there is a record of a dhole that interbred with a golden jackal. DNA sequencing of the Sardinian dhole (Cynotherium sardous) an extinct small canine species formerly native to the island of Sardinia in the Mediterranean, and which has often been suggested to have descended from Xenocyon, has found that it is most closely related to the living dhole among canines. Admixture with the African wild dog In 2018, whole genome sequencing was used to compare all members (apart from the black-backed and side-striped jackals) of the genus Canis, along with the dhole and the African wild dog (Lycaon pictus). There was strong evidence of ancient genetic admixture between the dhole and the African wild dog. Today, their ranges are remote from each other; however, during the Pleistocene era the dhole could be found as far west as Europe. The study proposes that the dhole's distribution may have once included the Middle East, from where it may have admixed with the African wild dog in North Africa. However, there is no evidence of the dhole having existed in the Middle East nor North Africa, though the Lycaon was present in Europe during the Early Pleistocene, with its last record in the region dating to 830,000 years ago. Genetic evidence from the Sardinan dhole suggests that both Sardinian and modern dholes (which are estimated to have split from each other around 900,000 years ago) share ancestry from the Lycaon lineage, but this ancestry is significantly higher in modern dholes than in the Sardinian dhole. , seven subspecies are recognised. In the absence of further data, the researchers involved in the study speculated that Javan and Sumatran dholes could have been introduced to the islands by humans. Fossils of dhole from the early Middle Pleistocene have been found in Java. ==Characteristics==

, India (1890) The general tone of the dhole's fur is reddish, with the brightest hues appearing in winter. During the winter coat phase, the back is covered in a saturated rusty-red to reddish color, featuring brownish highlights along the top of the head, neck, and shoulders. The throat, chest, flanks, belly, and upper parts of the limbs are less vividly colored, exhibiting a more yellowish tone. The lower parts of the limbs are whitish, with dark brownish bands on the anterior sides of the forelimbs. The muzzle and forehead are greyish-reddish. The tail is very luxurious and fluffy, mainly a reddish-ocher color, with a dark brown tip. The summer coat is shorter, coarser, and darker. The dhole has a wide and massive skull with a well-developed sagittal crest, The rostrum is shorter than that of domestic dogs and most other canids. The upper molars are weak, being one third to one half the size of those of wolves and have only one cusp as opposed to between two and four, as is usual in canids, In appearance, the dhole has been variously described as combining the physical characteristics of the gray wolf and the red fox, and as being "cat-like" on account of its long backbone and slender limbs. ==Distribution and habitat==

, Maharashtra, India Historically, the dhole lived in Singapore and throughout Central Asia including Afghanistan, Kyrgyzstan, Kazakhstan, Mongolia, Tajikistan and Uzbekistan, though it is now considered to be regionally extinct in these regions. but it is now also extinct in South Korea, The current presence of dholes in North Korea and Pakistan is considered uncertain. This was the first record of dholes from the country in almost three decades. It possibly still lives in the Primorsky Krai province in far eastern Russia, where it was considered a rare and endangered species in 2004, with unconfirmed reports in the Pikthsa-Tigrovy Dom protected forest area; no sighting was reported in other areas since the late 1970s. Currently, no other recent reports are confirmed of dholes being present in Russia, so the IUCN considered them to be extinct in Russia. In 2011 to 2013, local government officials and herders reported the presence of several dhole packs at elevations of near Taxkorgan Nature Reserve in the Xinjiang Autonomous Region. Several packs and a female adult with pups were also recorded by camera traps at elevations of around in Yanchiwan National Nature Reserve in the northern Gansu Province in 2013–2014. Dholes have been also reported in the Altyn-Tagh Mountains. In China's Yunnan Province, dholes were recorded in Baima Xueshan Nature Reserve in 2010–2011. Dhole samples were obtained in Jiangxi Province in 2013. Confirmed records by camera-trapping since 2008 have occurred in southern and western Gansu province, southern Shaanxi province, southern Qinghai province, southern and western Yunnan province, western Sichuan province, the southern Xinjiang Autonomous Region and in the Southeastern Tibet Autonomous Region. There are also historical records of dhole dating to 1521–1935 in Hainan Island, but the species is no longer present and is estimated to have become extinct around 1942. Its presence was confirmed in the Kanchenjunga Conservation Area in 2011 by camera traps. In February 2020, dholes were sighted in the Vansda National Park, with camera traps confirming the presence of two individuals in May of the same year. This was the first confirmed sighting of dholes in Gujarat since 1970. In Bhutan, the dhole is present in Jigme Dorji National Park. In Bangladesh, it inhabits forest reserves in the Sylhet area, as well the Chittagong Hill Tracts in the southeast. Recent camera trap photos in the Chittagong in 2016 showed the continued presence of the dhole. These regions probably do not harbor a viable population, as mostly small groups or solitary individuals were sighted. Its range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam and Thailand, with the Vietnamese population considered to be possibly extinct. A camera trapping survey in the Khao Ang Rue Nai Wildlife Sanctuary in Thailand from January 2008 to February 2010 documented one healthy dhole pack. In northern Laos, dholes were studied in Nam Et-Phou Louey National Protected Area. In Vietnam, dholes were sighted only in Pu Mat National Park in 1999, in Yok Don National Park in 2003 and 2004; and in Ninh Thuan Province in 2014. A disjunct dhole population was reported in the area of Trabzon and Rize in northeastern Turkey near the border with Georgia in the 1990s. This report was not considered to be reliable. In August 2015, researchers from the National Museum of Natural History and the Karadeniz Technical University started an expedition to track and document possible Turkish population of dhole. In October 2015, they concluded that two skins of alleged dholes in Turkey probably belonged to dogs, pending DNA analysis of samples from the skins, and, having analysed photos of the skull of alleged dhole in Kabardino-Balkaria Republic of Russia, they concluded it was a grey wolf. ==Ecology and behaviour==

Dholes produce whistles resembling the calls of red foxes, sometimes rendered as coo-coo. How this sound is produced is unknown, though it is thought to help in coordinating the pack when travelling through thick brush. When attacking prey, they emit screaming KaKaKaKAA sounds. Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries. In contrast to wolves, dholes do not howl or bark. Social and territorial behaviour Dholes are more social than gray wolves, They live in clans rather than packs, as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of three to five animals, particularly during the spring season, as this is the optimal number for catching fawns. Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them. Clans typically number 5 to 12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals. and domestic dogs. They may defecate in conspicuous places, though a territorial function is unlikely, as faeces are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal latrines. They do not scrape the earth with their feet, as other canids do, to mark their territories. Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens. The gestation period lasts 60–63 days, with litter sizes averaging four to six pups. Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use rendezvous sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once weaning begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site for 70–80 days. By the age of six months, pups accompany the adults on hunts and will assist in killing large prey such as sambar by the age of eight months. Maximum longevity in captivity is 15–16 years. Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt at night, except on moonlit nights, indicating they greatly rely on sight when hunting. They can chase their prey for many hours. When chasing fleet-footed prey, they run at a pace of . (Rusa unicolor), Bandipur National Park Once large prey is caught, one dhole grabs the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat. They occasionally blind their prey by attacking the eyes. Serows are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes. Prey weighing less than is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes tear off pieces of the carcass and eat in seclusion. They give the pups access to a kill. They are generally tolerant of scavengers at their kills. Both mother and young are provided with regurgitated food by other pack members. There is one record of a pack bringing down an Indian elephant calf in Assam, despite desperate defense of the mother, resulting in numerous losses to the pack. Dholes eat fruit and vegetable matter more readily than other canids. In captivity, they eat various kinds of grasses, herbs and leaves, seemingly for pleasure rather than just when ill. In summertime in the Tian Shan Mountains, dholes eat large quantities of mountain rhubarb. Livestock predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. Oxen are killed more often than cows, probably because they are given less protection. Enemies and competitors , India In some areas, dholes are sympatric to tigers and leopards. Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap. Along with leopards, dholes typically target animals in the range (mean weights of for dhole and for leopard), while tigers selected for prey animals heavier than (but their mean prey weight was ). Also, other characteristics of the prey, such as sex, arboreality and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital, whereas leopards kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopards' greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight. Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike. Dhole packs are smaller in areas with higher tiger densities due to tigers directly killing dholes and stealing kills they made. The kleptoparasitism causes dholes to prefer hunting smaller animals because they can eat more of a smaller carcass before a tiger arrives to steal it. Direct predation can lead to lower reproductive and recruitment rates, lower hunting success rates and less food for the pups when a helper is killed, and potentially pack destabilization if one member of the breeding pair is killed. Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs. Dholes were once thought to be a major factor in reducing Asiatic cheetah populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes. Since leopards are smaller than tigers and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do towards tigers. Dhole packs occasionally attack Asiatic black bears, snow leopards and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves and lacerate their hindquarters. The dhole is also sympatric with the Indian wolf (Canis lupus pallipes) in parts of its range. There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary, and two observations in Satpura Tiger Reserve. They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion. Diseases and parasites Dholes are vulnerable to a number of different diseases, particularly in areas where they are sympatric with other canid species. Infectious pathogens such as Toxocara canis are present in their faeces. They may suffer from rabies, canine distemper, mange, trypanosomiasis, canine parvovirus and endoparasites such as cestodes and roundworms. ==Threats==

Habitat loss is thought to amount to 60% of the dhole's historical range in India. The fragmentation and isolation of dhole populations has resulted in inbreeding and the Allee effect, which threaten its long-term viability. Some ethnic groups like the Kuruba and Mon Khmer-speaking tribes will appropriate dhole kills; some Indian villagers welcome the dhole because of this appropriation of dhole kills. The Indochinese dhole population suffers heavily from nonselective hunting techniques such as snaring. The fur trade does not pose a significant threat to the dhole. The people of India do not eat dhole flesh and their fur is not considered overly valuable. Due to their rarity, dholes were never harvested for their skins in large numbers in the Soviet Union and were sometimes accepted as dog or wolf pelts (being labeled as "half wolf" for the latter). The winter fur was prized by the Chinese, who bought dhole pelts in Ussuriysk during the late 1860s for a few silver rubles. In the early 20th century, dhole pelts reached eight rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly. == Conservation ==

In India, the dhole is protected under Schedule 2 of the Wildlife Protection Act, 1972. The creation of reserves under Project Tiger provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the Indira Gandhi Zoological Park (IGZP) in Visakhapatnam. The dhole has been protected in Russia since 1974, though it is vulnerable to poison left out for wolves. In China, the animal is listed as a category II protected species under the Chinese wildlife protection act of 1988. In Cambodia, the dhole is protected from all hunting, while conservation laws in Vietnam limit extraction and utilisation. ==In culture and literature==

Three dhole-like animals are featured on the coping stone of the Bharhut stupa dating from 100 BC. They are shown waiting by a tree, with a woman or spirit trapped up it, a scene reminiscent of dholes treeing tigers. In Tamil Sangam literature from 100 BC to 200 AD, the dhole is a recurring character, a device of metonymy in song; Ainkurunuru 323 narrates a pregnant dhole craving to eat wild pig, so its male mate hunts for pig. Its fearsome reputation in India is reflected by the number of pejorative names it possesses in Hindi, which variously translate as "red devil", "devil dog", "jungle devil", or "hound of Kali". Leopold von Schrenck had trouble obtaining dhole specimens during his exploration of Amurland, as the local Gilyaks greatly feared the species. This fear and superstition was not shared by neighbouring Tungusic peoples. It was speculated that this differing attitude towards the dhole was due to the Tungusic peoples' more nomadic, hunter-gatherer lifestyle. Japanese author Uchida Roan wrote about the declining popularity of indigenous dog breeds in 1901, which he asserted were descended from the dhole. In China, the dhole was widely known throughout history and mythology. One notable legendary creature is the Nine sons of the dragon, which was believed to be a creature that was part-dhole, part-dragon. In modern times, the Chinese word for dhole is often confused with 'jackal' or 'wolf', resulting in many confusions and mistranslations of dholes as jackals or wolves. Tameability Brian Houghton Hodgson kept captured dholes in captivity, and found, with the exception of one animal, they remained shy and vicious even after 10 months. Adult dholes are nearly impossible to tame, though pups are docile and can even be allowed to play with domestic dog pups until they reach early adulthood. ==See also==